Thesis (PhD (Conservation Ecology and Entomology))--University of Stellenbosch, 2010. / ENGLISH SUMMARY: Climate change and biological invasions are major components of global change induced by human activity and are considered major drivers of global biodiversity decline in terrestrial ecosystems. These drivers interact synergistically and render ecosystems vulnerable to invasion by invasive alien species. Grasses are a group of invasive plants that easily respond to global changes and alter native plant community structure and ecosystem processes, such as fire frequency. To date there have been few studies that assess the ecological drivers and effects of invasive grass species on arid and semi-arid ecosystems of South Africa. Fountain grass (Pennisetum setaceum) is a widely distributed invasive alien perennial grass from North Africa and the Middle East, valued by horticulturalists worldwide. It spreads along the edges of roads on the outskirts of most towns, and is common on mine spoil in many areas throughout South Africa. Occasionally, it escapes into natural vegetation along drainage lines or after fires. This grass can be a costly problem for agriculture and biodiversity conservation as it is unpalatable and increases fire risk. Understanding its distribution and invasive dynamics will contribute to better management and control practices. The main objective of this study was to understand the ecology and invasion processes of P. setaceum across South African environmental gradients, and use it as a model to understand the synergistic relationships between biological invasions and other global change (climate and land use) scenarios. This study provides the first assessment of how P. setaceum overcomes different invasion barriers in South Africa as an emerging invader, in comparison with other parts of the world where it has already become problematic. A number of management and control options for this grass and other similar perennial grasses result from this study. Specifically, the study provides comprehensive understanding of: 1) the distribution and habitat preferences of P. setaceum in arid and semi-arid parts of South Africa, 2) environmental resources and habitat conditions that promote its invasive potential, 3) growth and reproductive performance across environmental gradients, 5) the response of an arid ecosystem to P. setaceum invasion and fire promotion, and 6) the predicted future distribution of this grass in South Africa under scenarios of climate change and spread. The study has found that road-river interchanges are ideal sites for P. setaceum growth and that these sites should be targeted for management and control of this species. P. setaceum was found to persist under diverse biome environments, which is attributed to local adaptation. Disturbance was found to be a major factor promoting fountain grass invasion into semi-natural areas away from roadsides. Major clean-ups of this grass should focus on disturbed areas, especially in the fertile parts of the fynbos region where the grass has high climatic suitability. Management and control should also focus on areas with high nutrients and extra water, as these areas facilitate growth and reproduction. The dynamics of P. setaceum invasion and spread makes it a good model for management of similar emerging invasive perennial grasses in similar ecosystems. / AFRIKAANSE OPSOMMING: Klimaatsverandering en biologiese indringing is grootskaalse komponente van mensgedrewe globale veranderinge, en kan ook gesien word as prominente drywers van die huidige afname in globale biodiversiteit van terrestriële ekosisteme. Hierdie drywers werk saam in sinergie, en laat sodoende ekosisteme kwesbaar vir indringing deur uitheemse indringerplante. Grasse is ‘n groep indringerplante wat maklik reageer ten opsigte van globale veranderinge en verander inheemse plantgemeenskapstruktuur en ekosisteem prosesse, soos byvoorbeeld veldbrand frekwensie. Tot op hede is daar wynig studies gedoen wat die ekologiese drywers en effekte van indringergras spesies in droë en semi-droë ekosisteme van Suid-Afrika assesseer. Pronkgras (Pennisetum setaceum) is ‘n wyd verspreide meerjarige uitheemse indringergras, oorspronklik vanaf Noord-Afrika en die Midde-Ooste, en word waardeer deur tuinboukundiges wêreldwyd. Dit versprei al langs padrande aan die buitewyke van meeste dorpe, en is algemeen op ou mynhope in verskeie dele van Suid-Afrika. Somtyds ontsnap hierdie gras langs dreineringskanale of na veldbrande, en beland so in die omringende natuurlike plantegroei. Hierdie gras kan ‘n duur probleem word vir landbou asook biodiversiteit bewaring omdat dit onsmaaklik is vir diere, en dit verhoog veldbrand risiko. ‘n Beter begrip van pronkgras verspreiding en indringingdinamika sal bydra tot verbeterde bestuur en kontrole praktyke. Die hoofdoel van hierdie studie was om die ekologie en indringing prosesse van P. setaceum regoor Suid-Afrikaanse omgewingsgradiente te begryp, en dit dan te gebruik as ‘n model om die sinergistiese verhoudings tussen biologiese indringing en ander globale veranderinge (klimaat en grondgebruik) te verstaan. Die studie verskaf die eerste assessering van hoe P. setaceum verskillende indringing hindernisse in Suid-Afrika oormeester as ‘n opkomende indringer, in vergelyking met ander dele van die wêreld waar dit al klaar problematies is. Hierdie studie verskaf uiteindelik ‘n aantal bestuur en kontrole opsies vir hierdie en ander soortgelyke meerjarige grasse. Die studie verskaf spesifiek ‘n deeglike verstaan van: 1) die verspreiding en habitat voorkeure van P. setaceum in droë en semi-droë areas in Suid-Afrika, 2) omgewingshulpbronne en habitat toestande wat hierdie plant se indringing potensiaal verhoog, 3) groei- en voortplantingsvertoning oor verskillende omgewingsgradiente, 4) die reaksie wat ‘n droë ekosisteem toon teenoor P. setaceum indringing en veldbrand verhoging, en 5) die toekomstige verspreiding van hierdie gras in Suid-Afrika onder voorspellings van klimaatsverandering en verspreiding. Die studie het gevind dat pad-rivier tussengange ideale liggings is vir P. setaceum vestiging, en dat hierdie liggings dus geteiken moet word vir bestuur en kontrole doeleindes. P. setaceum toon volharding onder diverse bioom omgewings, wat toegeskryf word aan plaaslike aanpassing. Daar is gevind dat versteuring ‘n groot faktor is in die verhoging van pronkgras indringing in semi-natuurlike areas weg van padskouers. Grootskaalse pronkgras uitroeiing projekte moet fokus op versteurde gebiede, veral in die vrugbare dele van die fynbos streek, waar hierdie gras hoë klimaatsgeskiktheid toon. Bestuur en kontrole programme moet ook fokus op areas met hoë nutriente inhoud en ekstra water, aangesien hierdie areas groei en voortplanting vergemaklik. Die dinamika van P. setaceum indringing en verspreiding maak dit ‘n goeie model vir die bestuur van soortgelyke opkomende meerjarige indringergrasse in soortgelyke ekosisteme.

Les aires de répartition des arbres pourraient être grandement affectées par le changement climatique. Les résultats d’analyses paléogéographiques ont montré que ces dernières se sont déjà déplacées avec les variations passées du climat. Ces études ont permis de déterminer la direction et la vitesse de migration des espèces, données utilisées actuellement pour générer des prédictions sur l’évolution de l’aire de répartition des espèces forestières en réponse au réchauffement du climat. Cependant, le contexte écologique dans lequel les arbres font face à ces changements est très différent par rapport aux changements climatiques passés : l'augmentation actuelle des températures est plus rapide, les surfaces susceptibles d’être colonisées sont occupées par des écosystèmes très différents et variés (forêts, surfaces agricoles, zones urbaines). Par conséquent, les arbres pourront-ils faire face à la rapidité des changements globaux actuels ? Auront-ils la capacité de migrer pour trouver des conditions plus favorables ou pourront-ils s’adapter et survivre à de nouvelles conditions environnementales ? Dans un premier temps, l’analyse de données historiques (plans d’aménagements de l’Office Nationale des Forêts et Inventaires Forestiers Espagnols) a permis de mettre en évidence des évènements de colonisation et d’extirpation et de quantifier la vitesse de migration de populations situées au cœur ou aux marges de leur aire de répartition. Une colonisation massive de Quercus ilex dans les dunes boisées atlantiques (limite Nord d’aire de répartition) a été mise en évidence au cours des 130 dernières années, confirmant les tendances prédites par les modèles. Cependant, les vitesses de colonisation de cette espèce restent bien inférieures aux déplacements de son bioclimat estimés à partir de modèles de niche. Les espèces localisées en limite Sud d’aire de répartition présentent des remontées altitudinales plus importantes que pour celles situées au cœur de leur aire. En conclusion, nos résultats montrent que les changements globaux ont déjà impactés la répartition des arbres malgré l’existence d’un décalage temporel entre les réponses migratoires des espèces forestières et le déplacement de leur bioclimat. Le stress hydrique est le facteur prépondérant pouvant expliquer le dépérissement des arbres dans un milieu dont la disponibilité en eau est limitée, notamment en marge chaude d’aire de répartition. Nous avons, par conséquent, étudié la résistance à la sécheresse et ses mécanismes chez les plusieurs espèces d’Angiospermes. Nos résultats montrent qu’un seuil de 90% d’embolie mène à des dommages physiologiques irréversibles de la plante et à la mort par déshydratation. Ce seuil est considérablement plus élevé que celui précédemment observé chez les conifères. L’étude du fonctionnement hydraulique d’espèces de chênes co-occurrentes nous a permis de montrer que la survie de Q. robur pourrait être menacée dans les forêts atlantiques dans un contexte de sécheresses de plus en plus intenses car il y subit des taux d’embolie native élevés. Au contraire, Q. ilex présente des taux d’embolie négligeables sur ce même site d’étude.Les vitesses réelles de migrations constituent des données empiriques essentielles qui nous renseignent sur les capacités migratoires effectives des arbres. Elles pourront être intégrées dans les modèles de répartition, tout comme les seuils d’embolie induisant la mort des arbres. / Tree distribution could be highly affected by climate change. Results of paleogeographic studies showed that tree distribution ranges have already shifted with past climate changes. These data are currently used to model the evolution of species distribution in response to global warming. However, the ecological context in which species have to cope with climate change is very different than the past one: the current increase of temperature occurs faster than the past global warming, the areas being likely colonized are covered by various ecosystems (forests, agricultural surfaces, urban areas). So will tree species be able to cope with the current global change? Will they be able to migrate to find more favourable conditions or to survive to drier environmental conditions? Firstly, the analysis of historical data (French National Forest Office and Spanish National Forest Inventories) allowed determining colonization and extirpation events, and quantifying migration rates of tree species populations situated at the edges or the core of their distribution range. We evidenced that Q. ilex has substantially colonized new areas at its northern margin during the last 130 years, confirming the model trends. However, the colonization rates of Q. ilex are much lower than the shift of its bioclimate predicted by bioclimatic models. Species located at their rear edge showed higher upward shifts than other species located at the core of their range. To conclude, our results showed that global change have already impacted tree distribution although a time-lag between forest species migration responses and their bioclimate shift. Water stress is the main factor explaining tree dieback when water is limited and so particularly at the warm limit of tree species distribution range. Therefore, we studied drought resistance and its mechanisms in angiosperm tree species. Our results showed that embolism threshold of 90% leads to irreversible damages and tree death by dehydration. This threshold is considerably higher than the observed in Conifers. The study of hydraulic functioning of co-occurring oaks showed that the survival of Q. robur could be threatened in the context of increasing drought in the Atlantic forests because of its functioning at high levels of embolism. On the contrary, Q. ilex presented negligible levels of embolism in the same study area. The migration rates form primordial empirical data that give us information about tree effective migration abilities. They could be integrated within vegetation distribution models as well as embolism thresholds leading to tree mortality.

Les poissons migrateurs amphihalins évoluent dans un environnement qui est perturbé, à l’échelle globale, par le changement climatique et à l’échelle régionale, par les pressions anthropiques telles que les barrages et la pollution des cours d’eau. Cette étude, menée à l’échelle continentale, avait pour objectif d’évaluer, de comprendre et de prédire les impacts de changements globaux sur la distribution de ce groupe. Afin de répondre à cette problématique, des modèles empiriques de distribution ont été construits pour chacune des 28 espèces étudiées. Leurs distributions ont été décrites au début et à la fin du XXème siècle, en termes de présence-absence et de classes d’abondance. Les variables explicatives potentielles étaient de type physique, climatique, biotique et anthropique. Dans un premier temps, il a été mis en évidence que la répartition vers 1900 des poissons migrateurs amphihalins était fortement contrainte par leur histoire biogéographique et par les conditions thermiques et hydrauliques de l’habitat continental. La projection de ces distributions, à la fin du XXIème siècle, dans le contexte du changement climatique, a montré la forte sensibilité de ce groupe aux modifications du climat. Bien que spécifiques à chaque espèce, les réponses ont pu être classées en trois catégories : expansion de l’aire de répartition, contraction de l’aire de répartition et peu ou pas de changement. Dans un deuxième temps, pour la première fois à l’échelle continentale, l’importance des grands barrages et de la densité de populations humaines pour expliquer la distribution actuelle de ces espèces a été démontrée. Les relations sont apparues complexes et dépendantes des caractéristiques écologiques des espèces, notamment de la position de leurs habitats de croissance et de reproduction le long du réseau hydrographique et de leur place au sein du réseau trophique. / Diadromous fishes live in an environment disturbed, at global scale, by climate change and at regional scale, by anthropogenic pressures such as dams and river pollution. This study, performed at continental scale, aimed to assess, understand and predict the impacts of global changes on the distribution of this group. To address this issue, empirical distribution models were built for each of the 28 species studied. Their distributions were described at the beginning and the end of the twentieth century, in terms of presence-absence and abundance. Different types of explanatory variables were used, i.e. physical, climatic, biotic and anthropogenic. On the one hand, it was demonstrated that the distribution of diadromous fishes around 1900 was constrained by their biogeographical history and by the thermal and hydraulic characteristics of the continental habitat. Projection of these distributions, at the end of the twenty-first century, under climate change, highlighted the sensitivity of this group to climatic modifications. Responses were species-specific but could be classified into three categories: expansion of the distribution range, contraction of the distribution range and little or no change. On the other hand, for the first time at continental scale, the importance of large dams and human population density to explain the current diadromous fish distribution was demonstrated. The relationships seemed complex and to be dependent on the species’ ecological characteristics, particularly the position of their essential habitats along the river network and their place within the food web.

Anthropogenic activities are increasing inorganic nitrogen (N) loadings to lakes in the northern hemisphere. In many boreal lakes phytoplankton are N limited, wherefore enhanced N input may affect the productivity of pelagic food webs. Simultaneously, global change causes increased inflows of terrestrial dissolved organic carbon (DOC) to boreal lakes. Between clear and humic lakes, whole lake primary and consumer production naturally differs. However, research is inconclusive as to what controls pelagic production in these lakes. Further, it is unclear how DOC affects the response of the pelagic food web to enhanced inorganic N availability. The overarching goal of this thesis was to study the effects of inorganic N and organic C for pelagic food webs in boreal lakes. In the thesis, I first identified the main drivers of pelagic production during summer in eight non-manipulated Swedish boreal lakes with naturally low or high DOC. Then I investigated how increased N availability affects the pelagic food chain, and how the response differs with DOC. Therefore, whole lake inorganic N fertilization experiments were conducted in six Swedish boreal lakes across a DOC gradient (low, medium, high) divided into three lake pairs (control, N enriched) with one reference and two impact years. In each lake, I also investigated the response of zooplankton growth using in situ mesocosm experiments excluding planktivores. I found that humic boreal lakes had lower phytoplankton production and biomass than clear water lakes. Further, phytoplankton community composition and food quality differed with DOC. However, high DOC did not reduce pelagic energy mobilization or zooplankton biomass, but promoted a higher dominance of cladoceran relative to copepod species. N addition clearly enhanced phytoplankton biomass and production in the experimental lakes. However, this stimulating N effect decreased with DOC as caused by light limitation. Further, the newly available phytoplankton energy derived from N addition was not efficiently transferred to zooplankton, which indicates a mismatch between producer energy supply and consumer energy use. Indeed, the mesocosm experiment revealed that decreased food quality of phytoplankton in response to N addition resulted in reduced food web performance, especially in clearer lakes. In humic lakes, zooplankton production and food web efficiency were clearly more resilient to N addition. In summary, my thesis suggests that any change in the landscape that enhances inorganic N availability will especially affect pelagic food webs in clear water lakes. In contrast, brownification will result in more lakes being resilient to eutrophication caused by enhanced N deposition.

<p>Since the pre-industrial age, the Earth has been warming at unparalleled rates, and this warming is changing climate and weather, creating a more extreme global hydrological cycle. In this dissertation, I explore how these changes to the hydrological cycle may act ecosystem and community level responses of terrestrial plants in the Midwestern United States. In this region, it is projected that mean annual precipitation (MAP) will increase, but precipitation will become more variable across and within seasons. Ecosystem structure and function are vulnerable to changes in hydrologic patterns, including changes in biogeochemical cycles, plant productivity, and plant community structure and function. In this dissertation, I explore how changes in precipitation will alter these processes using two field experiments, and I suggest potential hypotheses that could explain drought-induced community change.</p><p><br></p><p>In chapter 1, I explore how alterations to seasonal precipitation in the winter and summer act ecosystem and community processes in a temperate deciduous forest. Biogeochemical processes and plant communities are sensitive to changes in abiotic conditions, and these conditions will alter forest succession, particularly juvenile woody plant species. Using a fully factorial experiment, I manipulated winter snowfall and summer precipitation to create wet, dry, and control (ambient conditions) treatments and investigated how changes in seasonal precipitation would act mineralization rates, woody plant recruitment, and understory composition. I found that the effects of winter and summer precipitation on these processes acted independently of one another in this system, and the system was resistant to changes in mineralization rates and understory composition. Woody plant recruitment may be more sensitive to altered precipitation, as recruitment of at least one of the four species planted, Lindera benzoin, was impacted by changes in seasonal precipitation. Snow removal treatments reduced germination and increased summer precipitation decreased the relative growth rate of this species. In the short term, slight changes to woody plant recruitment may have little impact on long-term forest succession, but as these changes persist over longer periods of time, they could alter the direction of succession, which could lead to changes in the understory community composition and nutrient cycling.</p> <p><br></p><p>The second and third chapters explore the effects that drought intensification will have on terrestrial plant communities. Numerous studies have investigated the effects of individual droughts on ecosystem and community responses, but the effects that both the timing and duration of drought have on these responses remain largely unknown. To explore this gap in the literature, I conducted a eld experiment using rainout shelters to reduce growing season precipitation, creating dry periods that varied in length and timing. Drought can impact productivity and diversity in this system, and the timing in which the drought occurs influences these effects. Surprisingly, I found that the length of drought did not affect productivity or community composition.</p> <p><br></p><p>The final chapter introduces the Community Response to Extreme Drought framework CRED), which addresses the potential temporal progression of mechanisms and plant-plant interactions that may lead to community changes during and after a drought. The mechanisms for the temporal evolution of community-level drought responses are not fully understood, but plant-plant interactions, both competitive (-) and facilitative (+), are increasingly being recognized as important drivers of community compositional changes. The CRED framework provides hypotheses for the roles that plant-plant interactions have on drought-induced community change. CRED addresses how system-specific variables and the intensity of drought may influence the strength of plant-plant interactions over time, and ultimately the systems resistance and resilience to drought. </p><p><br></p><p>The results from this dissertation work have revealed that more research needs to be done to fully understand how changes in precipitation regimes and patterns will affect terrestrial ecosystems and plant communities. A better understanding of how ecosystems and communities respond to drought timing and length can help improve climate models and restoration strategies.</p>

<p>Relevant information is essential for finding solutions in Integrated Water Management (IWM). Complex water systems and a need for increasing integration of sectors, actors and scales in IWM require new methods for developing and managing such information. This thesis investigates the role of information within the IWM process, as well as the main challenges for development of representative, accessibleand harmonized information. Results show how information needs and the information production process for IWM may be systematized, and indicate a large potential for information system development for IWM. However, in order to reach the full potential, today’s limited and heterogeneous water information needs to become more comprehensive, transparent, interoperable, dynamic, scalable and openly accessible. Large pressures on water systems are found in coastal catchment areas that are unmonitored across the local to the global scale, indicating a large importance of these areas for nutrient and pollutant loading. The globally accessible runoff data from catchment areas that are rich in pressures from population, agriculture and general economic activity further exhibit a rapidly declining trend during recent years. Major water system changes may therefore pass unnoticed if analyzed on the basis of openly accessible runoff global data. Furthermore, large discrepancies are found between land cover databases, which may result in major uncertainties in quantification of water and evapotranspiration flows. Identified information challenges may be relatively easily overcome by making better use of available information, while other challenges such as development of consistent baselines of core data and a possible re-prioritization of water-environmental monitoring programs may be both difficult and costly.</p>

<p>The aim of this study is to investigate the components of religion and spirituality in the specific context of "Fryshuset" – a multicultural and glocal arena. Against the backdrop of contemporary researchers such as Heelas and Woodhead and their hypothesis about the "spiritual revolution" and a "new spirituality", as well as the postmodern discussion about "the return of God", this study discusses how religious change can be understood in the context of Fryshuset.</p><p>The theoretical point of departure is taken in a hermeneutical method where Fleck´s theory about "thought-collective" and Gidden´s theory about the "duality of structure" are used in order to understand and describe the dialectical relationship between individual and social structure.</p><p>The method used in this study is qualitative. The interviews are made of six individuals representing different social projects at Fryshuset.</p><p>By analyzing the material from the field study through theories on religious change, three areas are identified:</p><ol><li>"The subjective turn"</li><li>Religion and spirituality</li><li>A Cosmopolitan interpretation of life </li></ol><p>The result of this study is that the religiosity among the interviewed can be described through Taylor’s understanding of the "subjective turn". Furthermore they are also part of a "double subjective turn" understood as an internalized subjective turn. Despite the fact that a lot of what the informants describe corresponds to Heelas and Woodheads theories of "subjective life spirituality" it is not relevant to talk about a "spiritual revolution". However, according to Beckford´s theories about religion as a "cultural resource" it can be described as a kind of new spirituality which is used in order to act and think global. The individuals use religion and spirituality as a cultural resource in their formation of a larger cosmopolitan interpretation of life.</p>

<p>At present, carbon sequestration in terrestrial ecosystems slows the growth rate of atmospheric CO₂ concentrations, and thereby reduces the impact of anthropogenic fossil fuel emissions on the climate system. Changes in climate and land use affect terrestrial biosphere structure and functioning at present, and will likely impact on the terrestrial carbon balance during the coming decades - potentially providing a positive feedback to the climate system due to soil carbon releases under a warmer climate. Quantifying changes, and the associated uncertainties, in regional terrestrial carbon budgets resulting from these effects is relevant for the scientific understanding of the Earth system and for long-term climate mitigation strategies.</p> <p>A model describing the relevant processes that govern the terrestrial carbon cycle is a necessary tool to project regional carbon budgets into the future. This study (1) provides an extensive evaluation of the parameter-based uncertainty in model results of a leading terrestrial biosphere model, the Lund-Potsdam-Jena Dynamic Global Vegetation Model (LPJ-DGVM), against a range of observations and under climate change, thereby complementing existing studies on other aspects of model uncertainty; (2) evaluates different hypotheses to explain the age-related decline in forest growth, both from theoretical and experimental evidence, and introduces the most promising hypothesis into the model; (3) demonstrates how forest statistics can be successfully integrated with process-based modelling to provide long-term constraints on regional-scale forest carbon budget estimates for a European forest case-study; and (4) elucidates the combined effects of land-use and climate changes on the present-day and future terrestrial carbon balance over Europe for four illustrative scenarios - implemented by four general circulation models - using a comprehensive description of different land-use types within the framework of LPJ-DGVM.</p> <p>This study presents a way to assess and reduce uncertainty in process-based terrestrial carbon estimates on a regional scale. The results of this study demonstrate that simulated present-day land-atmosphere carbon fluxes are relatively well constrained, despite considerable uncertainty in modelled net primary production. Process-based terrestrial modelling and forest statistics are successfully combined to improve model-based estimates of vegetation carbon stocks and their change over time. Application of the advanced model for 77 European provinces shows that model-based estimates of biomass development with stand age compare favourably with forest inventory-based estimates for different tree species. Driven by historic changes in climate, atmospheric CO₂ concentration, forest area and wood demand between 1948 and 2000, the model predicts European-scale, present-day age structure of forests, ratio of biomass removals to increment, and vegetation carbon sequestration rates that are consistent with inventory-based estimates. Alternative scenarios of climate and land-use change in the 21^st century suggest carbon sequestration in the European terrestrial biosphere during the coming decades will likely be on magnitudes relevant to climate mitigation strategies. However, the uptake rates are small in comparison to the European emissions from fossil fuel combustion, and will likely decline towards the end of the century. Uncertainty in climate change projections is a key driver for uncertainty in simulated land-atmosphere carbon fluxes and needs to be accounted for in mitigation studies of the terrestrial biosphere.</p> / <p>Kohlenstoffspeicherung in terrestrischen Ökosystemen reduziert derzeit die Wirkung anthropogener CO₂-Emissionen auf das Klimasystem, indem sie die Wachstumsrate der atmosphärischer CO₂-Konzentration verlangsamt. Die heutige terrestrische Kohlenstoffbilanz wird wesentlich von Klima- und Landnutzungsänderungen beeinflusst. Diese Einflussfaktoren werden sich auch in den kommenden Dekaden auf die terrestrische Biosphäre auswirken, und dabei möglicherweise zu einer positiven Rückkopplung zwischen Biosphäre und Klimasystem aufgrund von starken Bodenkohlenstoffverlusten in einem wärmeren Klima führen. Quantitative Abschätzungen der Wirkung dieser Einflussfaktoren - sowie der mit ihnen verbundenen Unsicherheit - auf die terrestrische Kohlenstoffbilanz sind daher sowohl für das Verständnis des Erdsystems, als auch für eine langfristig angelegte Klimaschutzpolitik relevant.</p> <p>Um regionale Kohlenstoffbilanzen in die Zukunft zu projizieren, sind Modelle erforderlich, die die wesentlichen Prozesse des terrestrischen Kohlenstoffkreislaufes beschreiben. Die vorliegende Arbeit (1) analysiert die parameterbasierte Unsicherheit in Modellergebnissen eines der führenden globalen terrestrischen Ökosystemmodelle (LPJ-DGVM) im Vergleich mit unterschiedlichen ökosystemaren Messgrößen, sowie unter Klimawandelprojektionen, und erweitert damit bereits vorliegende Studien zu anderen Aspekten der Modelunsicherheit; (2) diskutiert unter theoretischen und experimentellen Aspekten verschiedene Hypothesen über die altersbedingte Abnahme des Waldwachstums, und implementiert die vielversprechenste Hypothese in das Model; (3) zeigt für eine europäische Fallstudie, wie Waldbestandsstatistiken erfolgreich für eine verbesserte Abschätzung von regionalen Kohlenstoffbilanzen in Wäldern durch prozessbasierten Modelle angewandt werden können; (4) untersucht die Auswirkung möglicher zukünftiger Klima- und Landnutzungsänderungen auf die europäische Kohlenstoffbilanz anhand von vier verschiedenen illustrativen Szenarien, jeweils unter Berücksichtigung von Klimawandelprojektionen vier verschiedener Klimamodelle. Eine erweiterte Version von LPJ-DGVM findet hierfür Anwendung, die eine umfassende Beschreibung der Hauptlandnutzungstypen beinhaltet. </p> <p>Die vorliegende Arbeit stellt einen Ansatz vor, um Unsicherheiten in der prozessbasierten Abschätzung von terrestrischen Kohlenstoffbilanzen auf regionaler Skala zu untersuchen und zu reduzieren. Die Ergebnisse dieser Arbeit zeigen, dass der Nettokohlenstoffaustausch zwischen terrestrischer Biosphäre und Atmosphäre unter heutigen klimatischen Bedingungen relativ sicher abgeschätzt werden kann, obwohl erhebliche Unsicherheit über die modelbasierte terrestrische Nettoprimärproduktion existiert. Prozessbasierte Modellierung und Waldbestandsstatistiken wurden erfolgreich kombiniert, um verbesserte Abschätzungen von regionalen Kohlenstoffvorräten und ihrer Änderung mit der Zeit zu ermöglichen. Die Anwendung des angepassten Modells in 77 europäischen Regionen zeigt, dass modellbasierte Abschätzungen des Biomasseaufwuchses in Wäldern weitgehend mit inventarbasierten Abschätzungen für verschiede Baumarten übereinstimmen. Unter Berücksichtigung von historischen Änderungen in Klima, atmosphärischem CO₂-Gehalt, Waldfläche und Holzernte (1948-2000) reproduziert das Model auf europäischer Ebene die heutigen, auf Bestandsstatistiken beruhenden, Abschätzungen von Waldaltersstruktur, das Verhältnis von Zuwachs und Entnahme von Biomasse, sowie die Speicherungsraten im Kohlenstoffspeicher der Vegetation. Alternative Szenarien von zukünftigen Landnutzungs- und Klimaänderungen legen nahe, dass die Kohlenstoffaufnahme der europäischen terrestrischen Biosphäre von relevanter Größenordnung für Klimaschutzstrategien sind. Die Speicherungsraten sind jedoch klein im Vergleich zu den absoluten europäischen CO₂-Emissionen, und nehmen zudem sehr wahrscheinlich gegen Ende des 21. Jahrhunderts ab. Unsicherheiten in Klimaprojektionen sind eine Hauptursache für die Unsicherheiten in den modellbasierten Abschätzungen des zukünftigen Nettokohlenstoffaustausches und müssen daher in Klimaschutzanalysen der terrestrischen Biosphäre berücksichtigt werden.</p>

About 24 % of the land surface in the northern hemisphere are underlayed by permafrost in various states. Permafrost aggradation occurs under special environmental conditions with overall low annual precipitation rates and very low mean annual temperatures. Because the general permafrost occurrence is mainly driven by large-scale climatic conditions, the distribution of permafrost deposits can be considered as an important climate indicator. The region with the most extensive continuous permafrost is Siberia. In northeast Siberia, the ice- and organic-rich permafrost deposits of the Ice Complex are widely distributed. These deposits consist mostly of silty to fine-grained sandy sediments that were accumulated during the Late Pleistocene in an extensive plain on the then subaerial Laptev Sea shelf. One important precondition for the Ice Complex sedimentation was, that the Laptev Sea shelf was not glaciated during the Late Pleistocene, resulting in a mostly continuous accumulation of permafrost sediments for at least this period. This shelf landscape became inundated and eroded in large parts by the Holocene marine transgression after the Last Glacial Maximum. Remnants of this landscape are preserved only in the present day coastal areas.<br><br> Because the Ice Complex deposits contain a wide variety of palaeo-environmental proxies, it is an excellent palaeo-climate archive for the Late Quaternary in the region. Furthermore, the ice-rich Ice Complex deposits are sensible to climatic change, i.e. climate warming. Because of the large-scale climatic changes at the transition from the Pleistocene to the Holocene, the Ice Complex was subject to extensive thermokarst processes since the Early Holocene.<br><br> Permafrost deposits are not only an environmental indicator, but also an important climate factor. Tundra wetlands, which have developed in environments with aggrading permafrost, are considered a net sink for carbon, as organic matter is stored in peat or is syn-sedimentary frozen with permafrost aggradation. Contrary, the Holocene thermokarst development resulted in permafrost degradation and thus the release of formerly stored organic carbon. Modern tundra wetlands are also considered an important source for the climate-driving gas methane, originating mainly from microbial activity in the seasonal active layer.<br><br> Most scenarios for future global climate development predict a strong warming trend especially in the Arctic. Consequently, for the understanding of how permafrost deposits will react and contribute to such scenarios, it is necessary to investigate and evaluate ice-rich permafrost deposits like the widespread Ice Complex as climate indicator and climate factor during the Late Quaternary. Such investigations are a pre-condition for the precise modelling of future developments in permafrost distribution and the influence of permafrost degradation on global climate.<br><br> The focus of this work, which was conducted within the frame of the multi-disciplinary joint German-Russian research projects "Laptev Sea 2000" (1998-2002) and "Dynamics of Permafrost" (2003-2005), was twofold. First, the possibilities of using remote sensing and terrain modelling techniques for the observation of periglacial landscapes in Northeast Siberia in their present state was evaluated and applied to key sites in the Laptev Sea coastal lowlands. The key sites were situated in the eastern Laptev Sea (Bykovsky Peninsula and Khorogor Valley) and the western Laptev Sea (Cape Mamontovy Klyk region). For this task, techniques using CORONA satellite imagery, Landsat-7 satellite imagery, and digital elevation models were developed for the mapping of periglacial structures, which are especially indicative of permafrost degradation. The major goals were to quantify the extent of permafrost degradation structures and their distribution in the investigated key areas, and to establish techniques, which can be used also for the investigation of other regions with thermokarst occurrence. Geographical information systems were employed for the mapping, the spatial analysis, and the enhancement of classification results by rule-based stratification. The results from the key sites show, that thermokarst, and related processes and structures, completely re-shaped the former accumulation plain to a strongly degraded landscape, which is characterised by extensive deep depressions and erosional remnants of the Late Pleistocene surface. As a results of this rapid process, which in large parts happened within a short period during the Early Holocene, the hydrological and sedimentological regime was completely changed on a large scale. These events resulted also in a release of large amounts of organic carbon. Thermokarst is now the major component in the modern periglacial landscapes in terms of spatial extent, but also in its influence on hydrology, sedimentation and the development of vegetation assemblages. Second, the possibilities of using remote sensing and terrain modelling as a supplementary tool for palaeo-environmental reconstructions in the investigated regions were explored. For this task additionally a comprehensive cryolithological field database was developed for the Bykovsky Peninsula and the Khorogor Valley, which contains previously published data from boreholes, outcrops sections, subsurface samples, and subsurface samples, as well as additional own field data. The period covered by this database is mainly the Late Pleistocene and the Holocene, but also the basal deposits of the sedimentary sequence, interpreted as Pliocene to Early Pleistocene, are contained. Remote sensing was applied for the observation of periglacial strucures, which then were successfully related to distinct landscape development stages or time intervals in the investigation area. Terrain modelling was used for providing a general context of the landscape development. Finally, a scheme was developed describing mainly the Late Quaternary landscape evolution in this area. A major finding was the possibility of connecting periglacial surface structures to distinct landscape development stages, and thus use them as additional palaeo-environmental indicator together with other proxies for area-related palaeo-environmental reconstructions. In the landscape evolution scheme, i.e. of the genesis of the Late Pleistocene Ice Complex and the Holocene thermokarst development, some new aspects are presented in terms of sediment source and general sedimentation conditions. This findings apply also for other sites in the Laptev Sea region. / Die vorliegende Arbeit wurde im Rahmen der multidisziplinären Deutsch-Russischen Verbundprojekte "Laptev See 2000" (1998-2002) und "Dynamik des Permafrost" (2003-2005) erstellt.<br> Etwa 24 % der Landoberfläche der Erde sind von Permafrost unterlagert. Die ausgedehntesten Permafrostgebiete befinden sich heute in Sibirien. In Nordostsibirien, das während der letzten Eiszeit nicht von Inlandeismassen bedeckt bedeckt war, lagerten sich während dieser Zeit mächtige eisreiche Permafrostsedimente ab. Die durch den nacheiszeitlichen Meeresspiegelanstieg um ca. 120 Meter nur noch in den heutigen Küstengebieten erhaltenen Ablagerungen sind zum Teil hervorragende Paläoklimaarchive, die verschiedenste fossile organische Überreste der Eiszeitlichen Fauna und Flora konserviert haben. Aber auch die Sedimente und das enthalten Grundeis enthalten Klimainformationen z.B. die aus Mineralogie, Ablagerungsmilieu oder geochemischer und isotopenchemischer Zusammensetzung gewonnen werden können.<br><br> Der hohe Eisgehalt in den Sedimenten führte mit Beginn der holozänen Warmzeit zur Bildung von Thermokarst und Thermo-Erosion, d.h. zu starken Zersetzungserscheinungen durch Auftauen und Erosion. Thermokarst beschreibt das Schmelzen des Grundeises und die gleichzeitig stattfindende tiefe Absenkung der betroffenen Landoberfläche. Thermokarst geht mit der Bildung von Thermokarstseen einher, deren Wasserkörper ein zusätzlicher Wärmespeicher ist und das Auftauen des darunter liegenden Permafrost verstärken kann. In Sibirien, aber auch anderen Regionen der Arktis, sind weite Gebiete von Thermokarst betroffen. Der Einfluss dieser klimabedingten großräumigen Landschaftsveränderungen in Permafrostgebieten auf den lokalen, regionalen und auch globalen Stoff- und Energiehaushalt ist bisher nur wenig untersucht. Die vorliegende Arbeit beschäftigt sich mit der Charakterisierung und Evolution von periglazialen Landschaften im nordsibirischen Laptevsee-Gebiet, die seit dem Beginn des Holozän von solchen klimatisch bedingten Veränderungen betroffen sind, und liefert damit ein Puzzleteil zum einen für die Rekonstruktion der Landschaft und Landschaftsentwicklung als auch Vorraussetzungen für das Verständnis der großräumig wirkenden geologischen und geomorphologischen Veränderungsprozesse. Die generellen Schwerpunkte, für die die vorliegende Arbeit Informationen liefert, sind die Charakterisierung von periglazialen Relief- und Oberflächentypen und die Bestimmung ihrer räumlichen Verbreitung, die Identifizierung und Quantifizierung einzelner geologischer und geomorphologischer Prozesse in diesen Landschaften, und die Rekonstruktion der Entwicklung periglazialer Landschaften im Spätquartär für Schlüsselgebiete im Küstengebiet der nordsibirischen Laptevsee.<br><br> Um diese generellen Schwerpunkte zu erreichen, werden verschiedene Einzelziele in der Arbeit verfolgt:<br><br> Die Entwicklung and Anwendung von Satellitenfernerkundungstechniken zur Analyse periglazialer Landschaften in Nordsibirien. Dazu werden hochauflösende Corona-Satellitendaten und multispektrale Landsat-7 Satellitendaten verwendet.<br> Die Untersuchung von Satellitenbildern, mit dem Schwerpunkt auf Oberflächen, die von der Zersetzung des eisreichen Permafrosts betroffen sind<br> Die Entwicklung von hochauflösenden digitalen Geländemodellen für die geomorphologische Analyse in zwei Schlüsselgebieten<br> Die räumliche Untersuchung der gewonnenen Daten mit Hilfe von geographischen Informationssystemen, mit einem Schwerpunkt auf Form, Verteilung und Außmaß von holozänem Thermokarst<br> Das Sammeln und Auswerten von Felddaten, mit Schwerpunkt auf Oberflächeneigenschaften periglazialer Landschaften und der Zusammensetzung der Permafrostablagerungen<br> Die Anwendung der gewonnenen Daten zur Unterstützung, Verbesserung und Ausweitung der lokal gewonnenen Felddaten und Paläoumweltrekonstruktionen, sowie die datengestützte Entwicklung von Vorstellungen zur Landschaftsgenese<br><br> Weite, Permafrost-dominierte Küstentiefländer der heutigen Laptevsee in Nordost-Sibirien sind durch die spätpleistozänen Ablagerungen des Eiskomplex aufgebaut. Diese zumeist schluffig bis mittelsandigen Ablagerungen sind durch einen sehr großen Eisgehalt in Form von verteiltem Grundeis und großer syngenetischer Eiskeile, sowie einem relativ hohen Anteil an organischen Resten gekennzeichnet. Mit Beginn der holozänen Klimaerwärmung kam es zur weitläufigen Bildung von Thermokarst.

The present thesis aims to introduce process-based model for species range dynamics that can be fitted to abundance data. For this purpose, the well-studied Proteaceae species of the South African Cape Floristic Region (CFR) offer a great data set to fit process-based models. These species are subject to wildflower harvesting and environmental threats like habitat loss and climate change. The general introduction of this thesis presents shortly the available models for species distribution modelling. Subsequently, it presents the feasibility of process-based modelling. Finally, it introduces the study system as well as the objectives and layout. In Chapter 1, I present the process-based model for range dynamics and a statistical framework to fit it to abundance distribution data. The model has a spatially-explicit demographic submodel (describing dispersal, reproduction, mortality and local extinction) and an observation submodel (describing imperfect detection of individuals). The demographic submodel links species-specific habitat models describing the suitable habitat and process-based demographic models that consider local dynamics and anemochoric seed dispersal between populations. After testing the fitting framework with simulated data, I applied it to eight Proteaceae species with different demographic properties. Moreover, I assess the role of two other demographic mechanisms: positive (Allee effects) and negative density-dependence. Results indicate that Allee effects and overcompensatory local dynamics (including chaotic behaviour) seem to be important for several species. Most parameter estimates quantitatively agreed with independent data. Hence, the presented approach seemed to suit the demand of investigating non-equilibrium scenarios involving wildflower harvesting (Chapter 2) and environmental change (Chapter 3). The Chapter 2 addresses the impacts of wildflower harvesting. The chapter includes a sensitivity analysis over multiple spatial scales and demographic properties (dispersal ability, strength of Allee effects, maximum reproductive rate, adult mortality, local extinction probability and carrying capacity). Subsequently, harvesting effects are investigated on real case study species. Plant response to harvesting showed abrupt threshold behavior. Species with short-distance seed dispersal, strong Allee effects, low maximum reproductive rate, high mortality and high local extinction are most affected by harvesting. Larger spatial scales benefit species response, but the thresholds become sharper. The three case study species supported very low to moderate harvesting rates. Summarizing, demographic knowledge about the study system and careful identification of the spatial scale of interest should guide harvesting assessments and conservation of exploited species. The sensitivity analysis’ results can be used to qualitatively assess harvesting impacts for poorly studied species. I investigated in Chapter 3 the consequences of past habitat loss, future climate change and their interaction on plant response. I use the species-specific estimates of the best model describing local dynamics obtained in Chapter 1. Both habitat loss and climate change had strong negative impacts on species dynamics. Climate change affected mainly range size and range filling due to habitat reductions and shifts combined with low colonization. Habitat loss affected mostly local abundances. The scenario with both habitat loss and climate change was the worst for most species. However, this impact was better than expected by simple summing of separate effects of habitat loss and climate change. This is explained by shifting ranges to areas less affected by humans. Range size response was well predicted by the strength of environmental change, whereas range filling and local abundance responses were better explained by demographic properties. Hence, risk assessments under global change should consider demographic properties. Most surviving populations were restricted to refugia, serving as key conservation focus.The findings obtained for the study system as well as the advantages, limitations and potentials of the model presented here are further discussed in the General Discussion. In summary, the results indicate that 1) process-based demographic models for range dynamics can be fitted to data; 2) demographic processes improve species distribution models; 3) different species are subject to different processes and respond differently to environmental change and exploitation; 4) density regulation type and Allee effects should be considered when investigating range dynamics of species; 5) the consequences of wildflower harvesting, habitat loss and climate change could be disastrous for some species, but impacts vary depending on demographic properties; 6) wildflower harvesting impacts varies over spatial scale; 7) The effects of habitat loss and climate change are not always additive. / Das Ziel dieser Studie bestand daher darin, prozess-basierte Modelle zu entwickeln, die mit Daten zur Abundanz von Arten parametrisiert werden können. Die außergewöhnlich gut erforschten Proteaceen der südafrikanischen Kapregion (CFR), für die ein umfangreicher Datensatz zur Verfügung steht, stellen ein sehr geeignetes Untersuchungssystem zur Erstellung derartiger prozess-basierter Modelle dar. In Kapitel 1 beschreibe ich ein prozess-basiertes Modell für die Verbreitungsdynamik sowie die Methoden zur Parametrisierung des Modells mit Daten zu Abundanzverteilungen. Das Modell umfasst ein räumlich-explizites demographisches Modul und ein Beobachtungsmodul. Das demographische Modul verbindet artspezifische Habitatmodelle, die das geeignete Habitat beschreiben, und prozess-basierte demographische Modelle, die die lokale Dynamik und die Windausbreitung von Samen umfassen. Nach der Überprüfung der Parametrisierungs¬methoden mit simulierten Daten, wende ich die Modelle auf acht Proteaceenarten mit unterschiedlichen demographischen Eigenschaften an. Außerdem untersuche ich die Rolle von positiver (Allee-Effekte) und negativer Dichte-Abhängigkeit. Die Ergebnisse zeigen, dass Allee-Effekte und überkompensatorische Dynamik für viele Arten tatsächlich eine Rolle spielen. Der Großteil der geschätzten Parameter stimmt quantitativ mit unabhängigen Daten und beschreibt erfolgreich, wie die Abundanzverteilung aus der Bewegung und Interaktion der Individuen entsteht. Die vorgestellten Methoden scheinen daher zur Untersuchung von Ungleichgewichtsszenarien geeignet, die die Ernte von Infloreszenzen in Wildbeständen (Kapitel 2) und Umweltwandel (Kapitel 3) einschließen. In Kapitel 2 untersuche ich die Effekte der Ernte von Infloreszenzen in Wildbeständen. Das Kapitel beinhaltet eine Sensitivitätsanalyse über mehrere räumliche Skalen sowie demographische Eigenschaften. Darauf folgend wurden die Effekte der Ernte anhand von drei realen Arten untersucht. Die Reaktion der Pflanzen auf die Ernte zeigte ein Verhalten mit abrupten Schwellenwerten. Die durch die Ernte am stärksten gefährdeten Arten zeichneten sich durch kurze Samenausbreitungsdistanzen, starke Allee Effekte, geringe maximale Reproduktionsrate, hohe Mortalität und hohe lokale Aussterbewahrscheinlichkeit aus. Die Betrachtung größerer räumlicher Skalen wirkte sich trotz schärferer Grenzwerte positiv auf die Reaktion der Arten aus. Die drei untersuchten realen Arten konnten sehr geringe bis mittlere nachhaltige Ernteraten ertragen. Zusammenfassend lässt sich sagen, dass Kenntnisse über die Demographie des Untersuchungssystems und die umsichtige Identifizierung der zu betrachtenden räumlichen Skala zu einer besseren Einschätzung der Ernteintensität und der Naturschutzziele führen sollten. In Kapitel 3 wird die Reaktion der Arten auf vergangene Habitatverluste und zukünftigen Klimawandel sowie die Interaktion der beiden untersucht. Der Klimawandel wirkte sich dabei vornehmlich negativ auf die Größe des Verbreitungsgebiets und die Ausnutzung des potentiellen Habitats (‚Range Filling’) aus, wobei es zu einer Verschiebung des Habitats ohne erfolgreiche Kolonisierung kam. Der Habitatverlust reduzierte vor allem die lokalen Abundanzen. Die meisten Arten wurden vor allem durch das Szenario mit beiden Klimawandel und Habitatsverlust stark beeinträchtigt. Der negative Effekt war allerdings geringer als nach einer einfachen Aufsummierung der Einzeleffekte zu erwarten wäre. Dies erklärt sich aus einer Verschiebung des Verbreitungsgebiets der Arten in Regionen, in denen es in der Vergangenheit zu geringeren Habitatverlusten kam. Die Größe des Verbreitungsgebiets wurde am besten durch die Stärke des Umweltwandels vorhergesagt, wogegen das Range Filling und die lokalen Abundanzen hauptsächlich von den demographischen Eigenschaften abhingen. Aus diesen Ergebnissen lässt sich schließen, dass Abschätzungen des Aussterbensrisikos unter Umweltwandel demographische Eigenschaften einbeziehen sollten. Die meisten überlebenden Populationen waren auf Refugien reduziert, die im Fokus der Naturschutzmaßnahmen stehen sollten. Zusammenfassend zeigen die Ergebnisse, dass 1) prozess-basierte demographische Modelle für die Verbreitungsdynamik von Arten mit Daten parametrisierbar sind; 2) die Einbeziehung demographischer Prozesse die Modelle für die Verbreitung von Arten verbessert; 3) verschiedene Arten von unterschiedlichen Prozessen beeinflusst werden und unterschiedlich auf Umweltwandel und Beerntung reagieren; 4) Dichteregulierung und Allee-Effekte bei der Untersuchung der Verbreitungsdynamik von Arten berücksichtigt werden sollten; 5) die Ernte von Infloreszenzen in Wildbeständen, sowie Habitatverlust und Klimawandel für manche Arten katastrophale Folgen haben können, deren Effekte aber von den demographischen Eigenschaften abhängen; 6) der Einfluss der Beerntung in Abhängigkeit von der betrachteten räumlichen Skala variiert; 7) die Effekte von Habitatverlust und Klimawandel nicht additiv sind.